BFPP/Bull. Fr. Pêche Piscic.

(2006) 383 : 23-40 — 23 —

ANNUAL ACTIVITY OF THE NOBLE CRAYFISH (ASTACUS ASTACUS)

IN THE ORLJAVA RIVER (CROATIA)

FALLER M. (1), MAGUIRE I. (2), KLOBUČAR G. (2)

(1) K. Krešimira 34, 34000 Požega, Croatia, matejfaller@gmail.com

(2) Department of Biology, Faculty of Science, University of Zagreb, Rooseveltov trg 6,
10000 Zagreb, Croatia, E-Mail: imaguire@biol.pmf.hr, gklobuca@zg.biol.pmf.hr
Reçu le 16 juin 2005 Received June 16, 2006
Accepté le 5 décembre 2006 Accepté le 5 décembre 2006

ABSTRACT
We studied the annual activity of the noble crayfish (Astacus astacus) at three
sites along the Orljava River, in the continental part of Croatia, between August 2003 and
September 2004. Each site represented the typical characteristics of the upper, middle
and lower section of the river (5, 24 and 37 km from the spring, respectively). The biggest
population size was recorded on the most upstream site, with greatest structural variability
of bottom, high biotic index, and the lowest mean water temperature. Males dominated
in catch during the whole research period (total sex ratio was 1.77 males: 1 female). The
number of caught crayfish fluctuated during the year and their activity was positively
correlated with the water temperature. The crayfish catch within the two downstream sites
was dramatically lower in the autumn 2004 then the year before. No obvious reason could
be found; therefore we concluded that this was probably result of natural fluctuations
in population. Males were significantly longer than females on all three sites. Males and
females had similar percentages of injuries, mainly on claws and antennae. Crayfish were
active during the whole year, even when water temperature was just 1°C. Phases of life
cycle (moulting, active cement glands, mating, hatchlings) occurred a month later in
our population than in the Northern Europe populations, probably as a consequence of
differences in the climate.
Key-words: Astacus astacus, life cycle, population dynamics, Croatia.

L’ACTIVITÉ ANNUELLE DE L’ÉCREVISSE À PIEDS ROUGES (ASTACUS ASTACUS L.)

DANS LA RIVIERE ORLJAVA (CROATIE)

RÉSUMÉ
Nous avons étudié l’activité annuelle de l’écrevisse à pieds rouges (Astacus astacus)
sur trois sites de la rivière Orljava, Croatie continentale. Les résultats présentés ici ont été
obtenus d’août 2003 à septembre 2004. Chacun des sites présente les caractéristiques
typiques des parties supérieure, moyenne et inférieure de la rivière (5, 24 et 37 km de
la source, respectivement). La population la plus importante a été enregistrée sur le
site le plus élevé, où le substrat est le plus diversifié, l’indice biotique le plus grand et
la température moyenne de l’eau la plus basse. Les mâles ont été prélevés dans les
nasses pendant l’étude (proportion de 1,77 mâles pour 1 femelle). Le nombre d’écrevisses
Article available at http://www.kmae-journal.org or http://dx.doi.org/10.1051/kmae:2006003
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 24 —

capturées a fluctué au cours de l’année et leur activité est positivement corrélée à la

température de l’eau. Le nombre d’écrevisses dans les sites en aval a diminué fortement
en 2004. Comme nous n’avons pas trouvé de raison évidente pour cette diminution,
nous pouvons l’attribuer à la fluctuation naturelle de la population. Les mâles capturés
étaient significativement plus grands que les femelles. Les blessures observées sont
principalement la perte de pinces, d’antennes et de pattes. Les écrevisses étaient actives
pendant l’année entière, même quand la température de l’eau était de 1°C. Les fonctions
physiologiques (la mue, le développement des glandes, la reproduction, l’éclosion des
juvéniles) sont apparues un mois plus tard que dans les populations de l’Europe du Nord,
probablement à cause des différences de climat.
Mots-clés: Astacus astacus, dynamique de la population, cycle annuel, Croatie.

INTRODUCTION
The noble crayfish (Astacus astacus) is one of four native European crayfish species
inhabiting Croatian freshwater habitats. It is native in the Sava River and the Drava River
basins, but, due to its economic value, it has also been introduced into some water bodies
in the Adriatic Sea Basin (MAGUIRE and GOTTSTEIN-MATOČEC, 2004). At present, the
noble crayfish is considered as a threatened species and is protected by Croatian Law
(NARODNE NOVINE, 70/05, 7/06).
The life cycle of A. astacus has been described by different authors (CUKERZIS,
1988; ACKEFORS, 1999). The most detailed information concerning the biology of the
noble crayfish exists for populations in the Northern Europe (SKURDAL and TAUGBØL,
2002). Noble crayfish mate in autumn, and the length of their life cycle depends on the
climate and the habitat in which they live. The breeding season begins with the temperature
decline in autumn. Testes and ovaries mature between July and September (LAHTI and
LINDQUIST, 1983). Mating and spawning is not fixed to a specific temperature or date,
but a higher temperature delays the onset of mating. Mating takes place in autumn, during
September and October (HUNER and LINDQVIST, 1986; WESTIN and GYDEMO, 1986)
and usually lasts for 2 to 3 weeks. Extrusion and fertilisation of eggs depends on the time
of mating and water temperature and occurs in period from a few hours to up to 6 weeks
after mating. The egg development time is also influenced by temperature and may last
8-9 months for the northern crayfish populations. A cold water period (< 5-6°C) seems
necessary to secure high egg and juvenile survival. According to CUKERZIS (1988), the
hatchlings of noble crayfish are 8.5 to 9 mm total length. After two moults, in the third
stage reaching 13-15 mm they look like true adults and they are adapted for solitary life.
Females reach sexual maturity at a size that ranges from 6.2 cm total length in
localities with earlier maturity or slow growth to 8.5 cm total length in localities with late
maturity or fast growth. Mean length of females with active cement glands stretches from
7.6 till 10.5 cm, depending on the population. This corresponds to an age of 3-5 years.
Proportion of mature females varies between different populations and years (6–97%).
Males become mature at a size of 6.0–7.0 cm total length (ABRAHAMSSON, 1966).

MATERIAL AND METHODS

Description of sites
The study was conducted at three sites along the Orljava River, the left tributary of
the Sava River, between August 2003 and September 2004. The river is 89 km long, and
the sites were chosen to represent the upper, the middle and the lower section of the river
(distance from a spring was 5, 24 and 37 km respectively) (Figure 1).
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 25 —

Figure 1
Position of study sites (black dots) on the Orljava River. The map was adapted
from original 1:250 000 map of Croatia. Sites 1, 2 and 3 correspond to upper,
middle and lower part of the river, respectively.

Figure 1
Localisation des sites d’étude (points noirs) sur la rivière Orljava. La carte a été
adaptée à partir d’une carte originale de la Croatie au 1 : 250 000. Les sites 1, 2 et
3 correspondent aux parties supérieure, moyenne et inférieure de la rivière.
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 26 —

Site 1 is a brook flowing through dense beech forest with alder trees along its
banks. There is no human activity upstream that could cause pollution of the water. Site
2 is a stream flowing through agricultural area, but has a 10 to 20 m wide zone of alder
and willow trees on the banks. Site 3 is a small river that flows through the middle of an
agricultural area, downstream from few small villages, with mostly willow trees on the
banks. Other characteristics of these sites are shown in Table I. In this area crayfish plague
or non-indigenous crayfish species have never been recorded (MAGUIRE and GOTTSTEIN-
MATOČEC, 2004) and there is no commercial exploitation of crayfish. Presence of crayfish
was recorded on all three sites during summer and autumn of 2003 and this research was
continued until September 2004.

Table I
Characteristics of study sites on the Orljava River.

Tableau I
Caractéristiques des sites d’étude sur la rivière Orljava.

Characteristic Site 1 Site 2 Site 3

Altitude [meters above sea level] 462 196 159
Length of study stretch [m] 115 125 140
Mean Width [m] 3.12 4.86 9.50
Mean Depth [cm] 17.9 50.5 61.4
Mean Velocity [ms-1] 0.33 0.57 0.19
Percentage of substrate < 1.6 cm 34 55 86
Percentage of substrate 1.6–15 cm 38 31 5
Percentage of substrate > 15 cm 28 14 9

Environmental parameters
Water temperature was measured on each site during each survey using
thermometer and Gemini Data Loggers. These temperature values were used for the
analyses of differences in crayfish activity during the year. Data on the other physical and
chemical parameters of water were obtained from Croatian Waters (Public company for
general water management control, survey and informing upon water conditions). Their
sampling site is two kilometres downstream from the Site 3, so we considered their data
representative for the analyses of life conditions on that site. Data were collected once
a month during the whole period of our research. Following parameters were analysed:
water temperature, pH, conductivity, alkalinity, concentration of O2, oxygen saturation,
concentration of NH3, nitrites, nitrates, total, inorganic and organic nitrogen and total
phosphorus (Table II). Observed difference in the water temperature, in the August 2003,
measured by Croatian Waters (29°C) and us (16°C) must have been a consequence of
probably inappropriate measurement (in the shallow, isolated part of the river) done by
Croatian Waters and it did not reflect the real water temperature of the river so we have
excluded it from our analyses.
Quality samples of macrozoobenthos were also taken using kick sampling net and
biotic index (WOODIWISS, 1964) was calculated for each site. Values of biotic index were
10, 11, and 10 for the sites 1, 2 and 3, respectively.
Information on presence of fish was obtained from local fishermen; also, occasionally,
fish were caught in traps. On the Site 1, the only fish species recorded was bullhead,
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 27 —

Table II
Physico-chemical parameters of water measured monthly by Croatian Waters
near Site 3 from August 2003 till September 2004 (Valid N = 14).

Tableau II
Paramètres physiques et chimiques mesurés chaque mois, entre Août et
Septembre 2004, par Croatian Waters (N = 14).

Water Parameter Mean Min. Max. Std. Dev.

Temperature [°C] 14.43 1 29 8.75
pH 7.83 7.23 8.34 0.29
Conductivity [µS cm-1] 372.07 239 458 65.83
Alkalinity [mg CaCO3 dm-3] 163.73 105 195 29.27
Oxygen [mg O2 dm-3] 10.58 7.95 13.08 1.56
Oxygen saturation [%] 98.00 82.38 112.95 6.84
NH3 [mg N dm-3] 0.074 0.012 0.436 0.116
-
NO2 [mg N dm-3] 0.008 0.002 0.017 0.004
NO3- [mg N dm-3] 0.831 0.231 1.667 0.422
Total nitrogen [mg N dm-3] 1.632 0.970 2.921 0.570
Inorganic nitrogen [mg N dm-3] 0.915 0.250 1.893 0.473
Organic nitrogen [mg N dm-3] 0.716 0.318 1.564 0.380
Total phosphorus [mg P dm-3] 0.118 0.046 0.297 0.081

Cottus gobio. On two lower sites, much higher fish abundance was observed. On the
Site 2, brown trout, Salmo trutta fario and on the Site 3, chub, Leuciscus spp. were
occasionally caught in traps.

Crayfish trapping and processing

We used LiNi traps (WESTMAN et al., 1978) and small, hand made ones (from PVC
bottles with two funnel entrances made from plastic net mesh size = 1 mm). Both trap
types were baited with (0.5 cm thick) sausage slices. Crayfish were captured intensively
during August and September 2003, and from October 2003 to September 2004 once or
twice a month, during two nights. On sites 1, 2 and 3, we used 9, 11 and 14 LiNi traps
respectively. Hand-made traps were used only on the Site 1 (13 traps) because river is too
deep on the two lower sites to allow successful use of this trap type. Traps were placed
near banks, alternately, in 10 m intervals.
Each individual was sexed, cephalothoraxes length (CL) and total length (TL) were
measured. Symptoms of diseases and some physiological characteristics (moulting
stage, females with active cement glands and external eggs and crayfish with remains of
spermatophores attached to the abdomen) were also recorded. Epibionts were collected
once from live crayfish and determined in the laboratory as Branchiobdella parasita and/or
B. pentodonta. They were found on a great number of crayfish (93%, 92% and 91% of
crayfish on sites 1, 2 and 3, respectively). Only one individual (0.1% of population) with
signs of burn spot disease was found on the Site 1.
Crayfish population size was expressed as Catch Per Unit of Effort (CPUE), which
represents number of caught crayfish per trap per night. Comparison of annual activity
and CPUE between sites is made only with data for crayfish caught in LiNi traps, because
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 28 —

Table III
Total number of crayfish caught in LiNi traps and number of trapping nights
through the whole period of research on each site.

Tableau III
Nombre total d’écrevisses capturées avec des nasses LiNi et nombre de nuits de
capture pendant toute l’étude pour chaque site.

Site 1 Site 2 Site 3

Month Sex Total N min/mean std. Total N min/mean std. Total N min/mean std.
Year (nights) /max for N dev. (nights) /max for N dev. (nights) /max for N dev.
8 M 172(10) 11/17.20/23 4.05 50(8) 4/6.25/9 1.58 - - -
2003 F 139(10) 8/13.90/20 3.75 33(8) 3/4.13/7 1.25 - - -
9 M - - - 60(7) 1/4.57/13 3.99 44(4) 2/11.00/20 7.75
2003 F - - - 6(7) 0/0.86/2 0.90 16(4) 0/4.00/6 2.83
10 M 6(3) 0/2.00/4 2.00 17(3) 0/5.67/13 6.66 43(6) 4/7.17/13 3.54
2003 F 5(3) 0/1.67/4 2.08 9(3) 2/3.00/4 1.00 27(6) 3/4.50/8 2.35
11 M 6(4) 0/1.50/6 3.00 31(4) 4/7.75/11 3.30 25(4) 2/6.25/9 3.40
2003 F 0(4) 0/0/0 0 1(4) 0/0.25/1 0.50 0(4) 0/0/0 0
12 M 0(2) 0/0/0 0 0(2) 0/0/0 0 2(2) 1/1.00/1 0
2003 F 0(2) 0/0/0 0 0(2) 0/0/0 0 0(2) 0/0/0 0
1 M 1(2) 0/0.50/1 0.71 0(2) 0/0/0 0 0(2) 0/0/0 0
2004 F 0(2) 0/0/0 0 0(2) 0/0/0 0 0(2) 0/0/0 0
2 M 0(2) 0/0/0 0 0(2) 0/0/0 0 1(2) 0/0.50/1 0.71
2004 F 0(2) 0/0/0 0 0(2) 0/0/0 0 0(2) 0/0/0 0
3 M 0(2) 0/0/0 0 3(2) 1/1.50/2 0.71 3(2) 1/1.50/2 0.71
2004 F 0(2) 0/0/0 0 1(2) 0/0.50/1 0.71 0(2) 0/0/0 0
5 M 14(3) 2/4.67/6 2.31 2(3) 0/0.67/1 0.58 2(3) 0/0.67/1 0.58
2004 F 1(3) 0/0.33/1 0.58 0(3) 0/0/0 0 0(3) 0/0/0 0
6 M 5(2) 1/2.50/4 2.12 0(2) 0/0/0 0 0(2) 0/0/0 0
2004 F 1(2) 0/0.50/1 0.71 0(2) 0/0/0 0 0(2) 0/0/0 0
7 M - - - - - - 1(1) 1/1.00/1 0
2004 F - - - - - - 1(1) 1/1.00/1 0
8 M 9(1) 9/9.00/9 0 3(1) 3/3.00/3 0 - - -
2004 F 8(1) 8/8.00/8 0 0(1) 0/0/0 0 - - -
9 M 17(2) 7/8.50/10 2.12 0(2) 0/0/0 0 2(2) 1/1.00/1 1.41
2004 F 3(2) 1/1.50/2 0.71 1(2) 0/0.50/1 0.71 0(2) 0/0/0 0
Total M 230(33) 166(38) 123(30)
F 157(33) 51(38) 44(30)

only they were used on all three sites. Data about crayfish caught in hand made traps on
the Site 1 are used as additional information on their life cycle. Traps were placed on the
same position during the whole research period.
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 29 —

Data analyses
All statistical analyses (Pearson correlations, descriptive statistics, ANOVA with
Tukey test) were performed using Statistica 5.0. Results were considered statistically
significant at p < 0.05. Graphs and tables were made using MS Office Excel 2002 and
Statistica 5.0.

RESULTS
During research period total of 935 (597 males and 338 females) crayfish were
caught in LiNi and hand made traps. Total sex ratios (males/females) of crayfish caught
in LiNi traps through the year were 1.46, 3.25 and 2.80 for sites 1, 2 and 3, respectively.
Statistically described data for crayfish caught in LiNi traps are shown in Table III.
Statistically described cephalothorax length is shown in Table IV. The largest male had
7.30 cm (TL = 13.69 cm) and the largest female 5.91 cm (TL = 11.78 cm) of cephalothorax
length. We recorded statistically significant difference (ANOVA) in cephalothorax length
between males and females on each site (Site 1 t = 5.48 p << 0.05, Site 2 t = 6.62 p <<
0.05, and Site 3 t = 4.86 p << 0.05).
Mean cephalothorax length, both for males and females, was the smallest on the
Site 1 and it increases downstream (Table IV). Males’size differed statistically significant
between sites 1 and 2 (t = 46.23, p << 0.05), and sites 1 and 3 (t = 62.75, p << 0.05).
Females’size was significantly different between sites 1 and 3 (t = 52.48, p << 0.05) and
also between sites 2 and 3 (t = 22.76, p << 0.05).
During our research we recorded injuries mainly on claws and antennae, but also
on legs, rostrum, cephalothorax and abdomen. Detailed data about injuries are shown in
Table V.
CPUE and water temperatures measured during each survey are described in Table
VI. CPUE was significantly positively correlated with the water temperature on the Site
1 (males r = 0.774, p = 0.005 and females r = 0.648, p = 0.032) (Figure 2). On the Site 2
(males r = 0.403, p = 0.195 and females r = 0.486, p = 0.113) and the Site 3 (males r =
0.326, p = 0.324 and females r = 0.465, p = 0.149) correlation was positive, but statistically
not significant (Figures 3 and 4).
Differences in crayfish activity (Figures 2-4) between months were analysed using
ANOVA. Only statistically significant p values for differences in CPUEs between months
are shown in Table VII.

Table IV
Length of cephalothorax of crayfish caught in LiNi traps on each site.

Tableau IV
Longueur des céphalothorax d’écrevisses capturées avec des nasses LiNi pour
chaque site.

Valid N Mean/cm Min./cm Max./cm Std. Dev.

Males on Site 1 159 4.84 3.38 7.21 0.79
Females on Site 1 106 4.35 3.34 5.91 0.55
Males on Site 2 70 5.60 3.30 7.12 0.80
Females on Site 2 19 4.44 3.29 5.45 0.68
Males on Site 3 69 5.68 3.99 7.30 0.75
Females on Site 3 23 5.20 4.52 5.72 0.31
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 30 —

Table V
Percentage of damages of different parts of crayfish body caught in LiNi traps for
each site.

Tableau V
Pourcentage d’animaux capturés blessés à différentes parties du corps pour
chaque site.

Site 1 Site 2 Site 3

Sex males females males females males females
Antennae % 13.11 11.74 11.24 6.25 8.20 6.82
Claws % 15.74 22.17 17.75 9.38 9.02 4.55
Legs % 2.95 3.04 3.55 1.56 2.46 0
Rostrum % 0.98 0.87 2.96 0 1.64 2.27
Cephalothorax % 0.98 0.43 1.18 0 0 2.27
Abdomen % 3.93 1.30 3.55 0 6.56 9.09

Table VI
Water temperature measured during trapping and CPUE of males and females for
all three sites.

Tableau VI
Température mesurée pendant la capture et CPUE pour les mâles et les femelles
pour chaque site.

Mean Min. Max. Std. Dev.

Temperature [°C], Site 1 9.16 0.1 17.0 6.02
Temperature [°C], Site 2 10.66 0.9 17.0 5.70
Temperature [°C], Site 3 10.42 1.5 15.8 5.27
CPUE males, Site 1 0.46 0 1.91 0.60
CPUE females, Site 1 0.26 0 1.54 0.50
CPUE males, Site 2 0.25 0 0.78 0.30
CPUE females, Site 2 0.07 0 0.38 0.12
CPUE males, Site 3 0.20 0 0.79 0.26
CPUE females, Site 3 0.06 0 0.32 0.12

The results of ANOVA (Table VII) indicated that CPUE (Figure 2) for males on the
Site 1 in late summer and autumn was higher in the year 2003 (August) compared to year
2004, but differences were not significant (August p = 0.704 and September p = 0.196).
Activity of males on the Site 1 was significantly higher in August 2003 compared to all the
months from October 2003 till June 2004 (p < 0.05). Activity of females on the Site 1 was
significantly higher in August 2003 compared to all the months from October 2003 till June
2004 (p < 0.05); also there was no significant difference recorded between August 2003
and August 2004 (p = 0.834).
On the two lower sites CPUE in the year 2004 was much lower then in the previous
year (Figures 3 and 4), but results of ANOVA indicated that the differences were mainly
insignificant (Table VII). On the Site 2 CPUE for females was significantly higher in August
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 31 —

Figure 2
Relation between CPUE and the water temperature (measured during survey) on
Site 1 (upper site).

Figure 2
Relation entre CPUE et température de l’eau (mesurée pendant l’étude) sur le site
1 (partie supérieure).

Figure 3
Relation between CPUE and the water temperature (measured during survey) on
Site 2 (middle site).

Figure 3
Relation entre CPUE et température de l’eau (mesurée pendant l’étude) sur le site
2 (partie moyenne).
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 32 —

Figure 4
Relation between CPUE and the water temperature (measured during survey) on
Site 3 (lower site).

Figure 4
Relation entre CPUE et température de l’eau (mesurée pendant l’étude) sur le site
3 (partie inférieure).

Table VII
Significant p values for differences in CPUE between months.

Tableau VII
Valeurs significatives de p pour les différences de CPUE entre les mois.

Year and Males Site 1 Females Site 1 Females Site 2 Females Site 3
Month August 2003 August 2003 August 2003 October 2003
2003 08 n.s. n.s. n.s. n.s.
2003 09 n.s. n.s. 0,00015 n.s.
2003 10 0,00032 0,00037 n.s. n.s.
2003 11 0,00021 0,00020 0,00018 0,03150
2003 12 0,00059 0,00071 0,00303 n.s.
2004 01 0,00059 0,00071 0,00303 n.s.
2004 02 0,00059 0,00071 0,00303 n.s.
2004 03 0,00059 0,00071 0,01228 n.s.
2004 05 0,00191 0,00023 0,00031 n.s.
2004 06 0,00304 0,00103 0,00303 n.s.
2004 08 n.s. n.s. n.s. n.s.
2004 09 n.s. 0,00236 0,01228 n.s.
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 33 —

2003 compared with September 2003, period from November 2003 till June 2004 and
September 2004 (p < 0.05). On the Site 3 only significant decrease was in the number of
females in November 2003 compared with October 2003 (p < 0.05).
Correlations between CPUE on the Site 3 and values of physical and chemical
parameters (measured monthly) of water were calculated. CPUE for males was significantly
positively correlated with total and organic nitrogen content (r = 0.80 p = 0.009, r = 0.91,
p = 0.001 respectively), and negatively with pH (r = -0.763, p = 0.017). CPUE for females
was significantly positively correlated with organic nitrogen content (r = 0.746, p = 0.021)
and negatively with conductivity of water (r = -0.68, p = 0.044).
Visible physiological stages of crayfish were recorded during the research period
and they are shown in Table VIII. Crayfish were active throughout the whole year. The
lowest temperature on all three sites was in January, and crayfish activity was recorded
even during that period. On the 5th of January 2004 one male was caught in the hand
made trap covered with ice, while water temperature was only 1°C.

Table VIII
Annual cycle of some physiological functions per months. X is a real finding, x is
presumption based on a real finding.

Tableau VIII
Apparition de quelques fonctions physiologiques le long de l’année pendant
l’étude. X représente un résultat confirmé, x une présomption.

Month 8 9 10 11 12 1 2 3 4 5 6 7 8 9
Year ‘03 ‘03 ‘03 ‘03 ‘03 ‘04 ‘04 ‘04 ‘04 ‘04 ‘04 ‘04 ‘04 ‘04
Activity X X X X X X X X x X X X X X
Moulting X X X X X X
乆乆 cement glands X X X
么乆 mating X
乆乆 external eggs x x x x x X X x
juveniles 1st stage x
juveniles 3rd stage X

Moulting was recorded for the first time in August in both years, and it continued till
November in the year 2003. During the research we sometimes found crayfish and their
exuvia in hand made traps.
Females with visible cement glands were recorded from September till November.
The smallest female with visible cement glands was caught on the 1st of October 2003 on
the Site 1 and it had cephalothorax length of 3.48 cm (TL = 7.13 cm). Mean cephalothorax
length of females with visible cement glands was 4.96 cm (TL = 9.86 cm).
Mating occurred in November. On the 10th of November 2003 on the Site 3 male and
female, with remains of spermatophores on their abdomens, were caught in LiNi trap.
Females with external eggs were caught during May and June. Only one female with
two juveniles (cephalothorax length of 4.41 cm) was caught, on the 1st of August 2004, on
the Site 1. Juveniles total length was 1.53 and 1.40 cm, and they were at the 3rd moulting
stage.
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 34 —

DISCUSSION AND CONCLUSION

Our data on the activity of Astacus astacus in the Orljava River are based on the
crayfish caught by traps and therefore provide information only on the trapped proportion
of the populations. Trappability (activity) is mainly connected with food searching (BROWN
and BOWLER, 1977), but might also fluctuate in connection to the other factors such as
water temperature, light conditions etc. (ABRAHAMSSON, 1983).
In our study, like in the study by BROWN and BOWLER (1977), there was
always higher number of trapped males than females regardless of the season (Table
III) (Figures 2-4). Equal number of males and females was caught only on the Site 1 in
October 2003 (m:f = 6:5) and August 2004 (m:f = 9:8). Similar results were presented by
LUCIĆ (2004); in her study on the population of the noble crayfish in the Vukovina Lake,
total sex ratio was 2.88:1 (m: f); from June till September number of males and females
was almost equal 1.15:1 (m: f), while during May and from October till January males
prevailed in the catch. More balanced sex ratio (m: f = 1.09:1) was found in the Velika
Paklenica Stream population (MAGUIRE et al., 2002). In their study, males were dominant
during July, and females during August and September. ACKEFORS (1999) divided a year
into two parts depending on females’activity and sex ratios; from November till June males
were dominant with sex ratio from 6:1 (m: f) to 16:1 (m: f), and from July till October sex
ratio was close to 1. From November till June only females without external eggs were
caught. It is known that lower number of females trapped during the egg bearing period
is a consequence of berried females being trap shy (ACKEFORS, 1999; SKURDAL and
TAUGBØL, 2002). In addition, our results on sex ratio could be explained with the BROWN
and BOWLER (1977) suggestion that all females “avoid” traps irrespective of reproductive
state, so sex ratio does not always represent the real situation within the population.
In general, on all three sites males were larger than females (Table IV). Cephalothorax
length differed significantly between the two sexes due to sexual dimorphism
(ABRAHAMSSON, 1966; SKURDAL and TAUGBØL, 2002). MAGUIRE et al. (2002) and
LUCIĆ (2004) also found statistically significant differences between males and females in
both cephalothorax and total length.
BOHL (1997) pointed out that the proportion of damaged crayfish in a population
could indicate existence of violation by predators. In our study the biggest proportion
of damaged crayfish was recorded on the Site 1 (Table V), where we haven’t recorded
presence of predatory fish (or any other predator), so we can exclude predation as a cause
of injuries. At the same time, population density on the Site 1 was higher than on the sites
2 and 3 (Table VI). Explanation for the high percentage of injuries (specially claw injuries)
could be in aggressive interference between crayfish, as it was shown by BOHL (1997)
who was studding a dense population of Austropotamobius pallipes. In our research,
we also observed that damaged claws were the most common injury both at males and
females (Table V), so we could assume that claw injuries were consequence of violent
behaviour between crayfish. Similar results were obtained by SKURDAL et al., 1988, who
found that the biggest proportion of chiliped loss was in dense populations where crayfish
were not exploited.
Other possible reasons of injuries were difficulties during moulting or rolling stones
during torrents. While in the study by LUCIĆ (2004), percentage of injured males was higher
than percentage of injured females (29.17% and 9.7%, respectively), in our study, overall
percentage of injured males and females was similar (36.35% and 33.43%, respectively).
It is known that crayfish presence (among other factors) is strongly dependent
on the structure of the stream bottom (NIEMI, 1977; BOHL, 1987; LAURENT, 1988;
REYNOLDS, 2002) and higher population density is expected where greater morphometric
and structural variability exists (BOHL, 1997). Composition and size of the bottom
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 35 —

substrate have the greatest structural variability on the Site 1 (Table I), and that is probably
the reason why this site had the highest population density.
Recorded physical and chemical parameters of water (Table II) were similar to
values recorded for the noble crayfish habitats (NIEMI, 1977; BOHL, 1987; ACKEFORS,
1999; MAGUIRE et al., 2002; LUCIĆ, 2004).
Since most of the information on a life cycle of noble crayfish is based on the
research made on the northern European populations, one of the aims of this research was
to investigate if Croatian populations of A. astacus follow the same pattern and time frame
of life cycle. However, some data on the Croatian noble crayfish population exist (Table IX).
Study made by LUCIĆ (2004) provides data for population of A. astacus in the Vukovina
Lake near Zagreb (continental part of Croatia), while population of A. astacus in the Velika
Paklenica Stream (in the Adriatic Sea drainage) was described by MAGUIRE et al. (2002).
In Table IX these data are presented in comparison to data gathered by ACKEFORS
(1999) for Swedish population, as well as with data collected during our research on the
population of the Orljava River.
Our results on the crayfish activity based on the number of trapped crayfish
(Table VIII) (Figures 2-4) showed that crayfish were active during the whole year, but as
expected, their activity was lower during winter period. Our record of active crayfish at
water temperature of 1°C is similar to findings for A. astacus in a small lake in Sweden

Table IX
Data on the occurrence of different phases of life cycle of Astacus astacus from
research performed by us in comparison to other authors: ACKEFORS (1999),
MAGUIRE et al. (2002) and LUCIĆ (2004). The results are presented with letters O
(Orljava River), S (Sweden), P (Paklenica Stream) and V (Vukovina Lake).

Tableau IX
Apparition tout au long de l’année des différentes phases du cycle annuel
d’Astacus astacus lors de notre étude et dans les études de : ACKEFORS (1999),
MAGUIRE et al. (2002) et LUCIĆ (2004). Les résultats sont présentés avec les
lettres O (Rivière Orljava), S (Suède), P (Fleuve Paklenica) et V (Lac Vukovina).

Month 8 9 10 11 12 1 2 3 4 5 6 7
PV PV V V V V P PV PV PV
Activity
O O O O O O O O O O O
SPV SPV SV V PV SPV
Moulting
O O O O
V SPV SV S
乆乆 Cement glands
O O O
S S
么乆 Mating
O
S S S S S SP SP SP SP
乆乆 External eggs
O O
P SP
Hatching
O
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 36 —

(ACKEFORS, 1999). He found a few active crayfish in January while water temperature
was just above 0°C. ŚMIETANA (2003) also found numerous narrow-clawed crayfish
(Astacus leptodactylus) in Poland, active throughout the whole winter period. Different
results were obtained by LUCIĆ (2004) and TROSCHEL et al. (1995). They found that
the lowest temperatures on which noble crayfish and stone crayfish were active were
4.5°C and 5°C, respectively. In our study crayfish activity was the highest from August
till October, probably as consequence of females becoming more active after releasing
the juveniles and both sexes preparing for mating. It is known that crayfish activity is
influenced by water temperature (KIVIVUORI, 1977; BOHL, 1987; MAGUIRE et al., 2002).
Higher temperatures can raise an animal’s locomotion and metabolic rates and it is likely
that the trappability is greater at higher temperatures (ACKEFORS, 1999). A positive
correlation between the water temperature and the number of animals caught was found
for Austropotamobius torrentium (TROSCHEL et al, 1995; MAGUIRE, 2002), as well as
for A. astacus (MAGUIRE et al., 2002; LUCIĆ, 2004). In our research, CPUE was also
positively correlated with the water temperature (Figures 2-4). However only on the Site 1
this correlation is statistically significant. Absence of significant correlations on sites 2 and
3 is probably consequence of general decline in number of crayfish trapped. After water
temperature increased in the year 2004, simply crayfish were scarce and there was not
many crayfish whose activity would increase and influence on significance of correlation.
In accordance with TROSCHEL et al. (1995) and MAGUIRE et al. (2002), we found that
the amount of activity in spring did not increase proportionally to the raise of the water
temperature, but it seems likely that the raise of temperature in early spring was an initiator
of their activity (Figures 2-4).
CPUE dramatically decreased on sites 2 and 3 in the year 2004, compared to the
year 2003 (Figures 3 and 4) (Table VII). Decrease was also observed on the Site 1, but not
with the same intensity (Figure 2). Declines in the population number are usually attributed
to: (1) the deterioration of water quality and habitat destruction, (2) direct competition
with non-native crayfish and (3) the crayfish plague (BRAMARD et al. 2003). None of
these reasons could explain the observed population decline in our study. DEMERS et
al. (2004) detected decline in the population of A. pallipes from Western France. In their
case, CPUE dropped to one tenth, and decline was explained as a consequence of low
oxygen concentration (average 6.2 mg/l) and high water temperature (maximum between
19°C and 29°C). Contrary to these findings, HOLDICH et al. (2005) has shown that A.
pallipes can also be found in muddy habitats, which are usually associated with low
oxygen concentrations. In our case, water temperatures, as well as oxygen concentrations
(minimum 8 mg l-1) were within the range that is suitable for noble crayfish (BOHL, 1987).
Thus decrease in population density on sites 2 and 3 cannot be explained by those
physical and chemical parameters. Normal fluctuations in crayfish density are also known
(REYNOLDS, 2002), so it is possible that recorded declines in our population density were
consequence of natural fluctuations in the population. As study on those populations is
still ongoing, we suppose that we will get more useful information about fluctuation in
population number that will help us explain declines.
According to the literature (SKURDAL and TAUGBØL, 2002), the first moulting period
for males and unberried females normally occurs in mid spring or early summer, depending
on the climate. The second moulting period occurs in late summer or early autumn, at
which time the females that were berried join in moulting. It is known that during moulting
animals are less active (REYNOLDS, 2002). We found freshly moulted crayfish only from
August to November (the second moulting period), since we have presumably missed the
first moulting period. ACKEFORS (1999) found moulting crayfish from July till October
what is one month less then in the Orljava River populations. Prolongation of moulting
period in this southern region could be explained with higher water temperature in autumn
and thus longer period of activity. MAGUIRE et al. (2002) found that moulting in coastal
stream populations started in June and lasted till September, while LUCIĆ (2004) found
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 37 —

that moulting in the lake population started even earlier, in May, and lasted till October
(Table IX). Finding of a freshly moulted crayfish inside the hand made trap indicates that
crayfish possibly used this trap type as a shelter.
Occurrence of cement glands clearly defines sexually mature females (GRANDJEAN
et al., 1997a, 1997b). The smallest female with developed cement glands that we caught
was 7.13 cm long and that is similar to findings of other authors (TAUGBØL et al., 1988;
WESTIN and GYDEMO, 1989; ACKEFORS, 1999). ACKEFORS (1999) found females with
active cement glands from September till November, and that is similar to results for our
population.
We recorded occurrence of mating in November. In Sweden mating occurred during
September and October (WESTIN and GYDEMO, 1986) and October and November
(ACKEFORS, 1999), while in Finland during September and October (HUNER and
LINDQVIST, 1986). It can be therefore concluded, that in Croatia mating starts later in
comparison to the Northern Europe, most probably due to prolonged period of higher
water temperatures in the autumn.
During our research we caught only two females with external eggs (in May and
June). Similarly, MAGUIRE et al. (2002) found external eggs till June, while LUCIĆ (2004)
did not find any female with external eggs. As said before, egg-bearing females are less
active, and therefore their catchability is lower so recorded percentages under represent
the actual number of berried females in the populations. In northern A. astacus populations
(ACKEFORS, 1999) females also carried eggs till June.
The appearance of juveniles in the 3rd stage was recorded on the 1st August 2004
(Table VIII) and that means that hatching probably occurred in July (Table IX). Eggs were
incubated for eight months, what is one month less then in the Swedish population studied
by ACKEFORS (1999) - hatchlings occurred in July, after nine months of incubation, and
similar to results from SKURDAL and TAUGBØL (2002) - the egg incubation period lasted
for 8-9 months.
The results of our research show that studied Croatian populations of A. astacus
follow the same pattern of life cycle as Northern European populations, but the time frame
is to some extent different probably due to the differences in the climate.

ACKNOWLEDGEMENT
We would like to thank the referees for their useful comments and suggestions that
helped to improve this paper.

REFERENCES
ABRAHAMSSON S.A.A., 1966. Dynamics of an isolated population of the crayfish Astacus
astacus Linné. Oikos, 17, 96-107.
ABRAHAMSSON S.A.A., 1983. Trappability, locomotion and diel pattern of activity of
crayfish A. astacus and P. leniusculus. Freshwater Crayfish, 5, 239-253.
ACKEFORS H.E.G., 1999. Observations on the Yearly Life Cycle of Astacus astacus in a
Small Lake in Sweden. Freshwater Crayfish, 12, 413-429.
BOHL E., 1987. Comparative studies on crayfish brooks in Bavaria (Astacus astacus L.,
Austropotamobius torrentium Schr.). Freshwater Crayfish, 7, 287-294.
BOHL E., 1997. An isolated population of the white-clawed crayfish (Austropotamobius
pallipes) in the principality of Liechtenstein. Bull. Fr. Pêche Piscic., 347, 701-712.
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 38 —

BRAMARD M., BACHELIER E., DUMAS J-C., FOURNIER C., BROUSSARD E., ROBIN O.,
GRANDJEAN F., 2003. Evolution of the distribution of native and exotic species
for twenty five years in Poitou-Charentes region (France) and causes of regression
of Autropotamobius pallipes. Abstract from The endangered native crayfish
Austropotamobius pallipes, Bioindicator and heritage species, 2, Craynet, Kilkenny,
Ireland.
BROWN D.J., BOWLER K., 1977. A population study of the British freshwater crayfish
Austropotamobius pallipes (Lereboullet). Freshwater Crayfish, 3, 33-49.
CUKERZIS, J.M., 1988. Astacus astacus in Europe. In: HOLDICH D.M., LOWERY R.S.
(eds.) Freshwater Crayfish: Biology, Management and Exploitation, 309-340,
Croom Helm, London.
DEMERS A., TROUILHE M-C., SOUTY-GROSSET C., BACHELIER E., GRANDJEAN F.,
2004. Impact of a heat wave on crayfish populations in a region of Western France.
Abstract from the European native crayfish in relation to land-use and habitat
deterioration, Craynet, Innsbruck, Austria.
GRANDJEAN F., ROMAIN D., AVILA-ZARZA C., BRAMARD M., SOUTY-GROSSET C.,
MOCQUARD J.P., 1997a. Morphometry, sexual dimorphism and size at maturity
of the white-clawed crayfish Austropotamobius pallipes pallipes (Lereboullet) from
a wild French population at Deux-Sèvres (Decapoda, Astacidae). Crustaceana, 70
(1), 31-44.
GRANDJEAN F., ROMAIN D., SOUTY-GROSSET C., MOCQUARD J.P., 1997b. Size at sexual
maturity and morphometric variability in three populations of Austropotamobius
pallipes pallipes (Lereboullet, 1858) according to a restocking strategy. Crustaceana,
70, 454-468.
HOLDICH D.M., FOSTER J., PEAY S., BRICKLAND J., 2005. Where does the white-clawed
crayfish live in muddy habitats? Abstract from the Craynet Final Conference, 20,
Craynet, Florence, Italy.
HUNER J.V., LINDQVIST O.V., 1986. A stunted crayfish Astacus astacus population in
central Finland. Freshwater Crayfish, 6, 156-165.
KIVIVUORI L., 1977. Temperature acclimation of the motor activity in the crayfish Astacus
astacus. Freshwater Crayfish, 4, 265-274.
LAHTI E., LINDQUIST O.V., 1983. On the reproductive cycle of the crayfish Astacus
astacus L. in Finland. Freshwater Crayfish, 5, 18-26.
LAURENT P.J., 1988. Austropotamobius pallipes and A. torrentium, with observations on
their interactions with other species in Europe. In HOLDICH D.M., LOWERY R.S.
(eds.), Freshwater crayfish: Biology, Management and Exploitation, 83-114, Croom
Helm, London.
LUCIĆ A., 2004. Physiological characteristics of three species of the freshwater crayfish
from the Astacidae family. PhD thesis (in Croatian with English abstract), 122 pages,
University of Zagreb, Croatia.
MAGUIRE I., 2002. Family Astacidae in north-west Croatia. PhD thesis (in Croatian with
English abstract), 128 pages, University of Zagreb, Croatia.
MAGUIRE I., ERBEN R., KLOBUČAR G.I.V., ŠTAMBUK A., HUDINA S., GALIĆ N., ÐURIĆ
P., 2002. Riječni rak (Astacus astacus) na području NP Paklenica. Study report for
NP Paklenica, 16 pages.
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 23-40 — 39 —

MAGUIRE I., GOTTSTEIN-MATOČEC S., 2004. The distribution pattern of freshwater

crayfish in Croatia. Crustaceana, 77 (1), 25-49.
NARODNE NOVINE 70, 2005. Zakon o zaštiti prirode.
NARODNE NOVINE 07, 2006. Pravilnik o proglašavanju divljih svojti zaštićenim i strogo
zaštićenim.
NIEMI A., 1977. Population studies on the crayfish Astacus astacus L. in the river Pyhäjoki,
Finland. Freshwater Crayfish, 3, 81-94.
REYNOLDS J.D., 2002. Growth and reproduction. In: HOLDICH D.M. (ed.), Biology of
Freshwater Crayfish, 152-192, Blackwell Science, Oxford.
SKURDAL J., TAUGBØL T., FJELD E., QVENILD T., 1988. Chiliped loss in Astacus astacus.
Freshwater Crayfish, 7, 47-51.
SKURDAL J., TAUGBØL T., 2002. Astacus. In: HOLDICH D.M. (ed.) Biology of Freshwater
Crayfish, 467-510, Blackwell Science, Oxford.
ŚMIETANA P., 2003. Migration and distribution of narrow-clawed crayfish Astacus
leptodactylus Esch. in the Lake Bylice, Wołczkowo. Oficyna In Plus: 106-110 (in
Polish with English abstract).
TAUGBØL T., SKURDAL J., FJELD E., 1988. Maturity and fecundity of Astacus astacus
females in Norway. Freshwater Crayfish, 7, 107-114.
TROSCHEL H.J., SCHULZ U., BERG R., 1995. Seasonal activity of stone crayfish
Austropotamobius torrentium. Freshwater Crayfish, 10, 196-199.
WESTIN L., GYDEMO R., 1986. Influence of light and temperature on reproduction and
moulting frequency in the crayfish, Astacus astacus. Aquaculture, 52, 43-50.
WESTIN L., GYDEMO R., 1989. Variation in sex ratio in the noble crayfish Astacus astacus:
reflection of the yearly activity changes. Freshwater Crayfish, 7, 115-120.
WESTMAN K., PURSIAINEN M., VILKMAN R., 1978. A new folding trap model which
prevents crayfish from escaping. Freshwater Crayfish, 4, 235-242.
WOODIWISS, F. S., 1964. The biological system of stream classification used by the Trent
River Board. Chemistry and Industry, 11, 443-447.
BFPP/Bull. Fr. Pêche Piscic. (2006) 383 : 41 — 41 —

REMERCIEMENTS :

Nous tenons à remercier chaleureusement l’ensemble des relecteurs ayant contribué

à la publication du BFPP pendant l’année 2006 :

Our warmest thanks to all the reviewers who have greatly contributed to the
publications of the BFPP during the year 2006:
Jan BAER, Vassilis BAKOPOULOS, Sanja BARIC, Roger BERGSTED, Eric BOHL,
Thierry BOUJARD, Christian BRY, José CARRAL, Daniel CATTAERT, Thomas CHANGEUX,
Edo D’AGARO, Andréanne DEMERS, Javier DIEGUEZ-URIDIBEONDO, Yves DESAUNAY,
Zdenek DÚRIS, Avi ELDAR, Sarah FRATINI, Leopold FÜREDER, Stuart R. GELDER,
Francesca GHERARDI, Piero GIULIANINI, Frédéric GRANDJEAN, Nicolas GOUIN,
Patrick HAFFNER, Paula HENTTONEN, David HOLDICH, Jay V. HUNER, Jussila JAPO,
Philippe KEITH, Max KELLER, Pavel KOZAK, Carlo R. LARGIARDÈR, Pierre LAURENT,
Marc LEGENDRE, Mario LEPAGE, Ivanna MAGUIRE, Ari MANNONEN, Charles MÉLARD,
Jean MORIN, Yann MOREAU, Pietro Angelo NARDI, Alix NIHOUARN, Pierre NÖEL, Birgit
OIDTMANN, Jean-Marc OLIVIER, Giuliana PARISI, Stephanie PEAY, Didier PONT, Jean-
Pierre PROTEAU, Barbara RENAI, Przemyslaw ŚMIETANA, Julian REYNOLDS, Charles
ROQUEPLO, Jean-Marc ROUSSEL, Maria SAEZ-ROYUELA, Massimiliano SCALICI,
Christoph SCHUBART, Holger SCHULZ, Ralf SCHULZ, Peter SIBLEY, Olivier SIMON, Fred
SLATER, Ondrej SLAVIK, Catherine SOUTY-GROSSET, Todd STEEVES, Thomas STUCKI,
Marc SUQUET, Evelyne TALÈS, Trond TAUGBØL, Ian WINFIELD, Serena ZACCARA.